After Conway Morris, 1993, and Conway Morris & Caron, 2007

Alternative phylogeny after Aguinaldo et. al. 1997

Alternative phylogeny after Winnepennickyx et al., 1998

Alternative phylogeny after Zrzavý et al., 2001

Alternative phylogeny after Brusca " Brusca, 2002

Alternative phylogeny after Paps et al., 2009

<==o Protostoma (protostomes; alkusuiset)
   |--o Aschelmithes [Pseudocoelomata] (valeruumiinonteloiset)
   |  |?- Tardigrada Spallanzani, 1777 (water bears; karhukaiset)
   |  |-- Chaetognatha Leuckart, 1854 [incl. †Protoconodonta] (arrow worms; nuolimadot)
   |  `--+--+-- Nematomorpha Vejdovsky, 1886 (horse hair worms, gordian worms; jouhimadot)
   |     |  `-- Nemata Gobb, 1919 [Nematoda Diesing, 1861] (nematodes, round worms; sukkulamadot)
   |     `--+-- Gastrotricha Metschnikoff, 1865 (hairybacks; sukaspintaiset)
   |        `--o Syndermata
   |           |== Rotifera Cuvier, 1798 (rotifers, wheel animals; rataseläimet)
   |           `-- Acanthocephala Koelreuter, 1771 (thorny-headed worms, spiny-headed worms; väkäkärsämadot)
   `--+?-+?- Entoprocta Nitsche, 1870 (entoprocts, goblet worms; pikarimadot)
      |  |?- Bryozoa Ehrenberg, 1831 [Ectoprocta Nitsche, 1869] (moss animals; sammaleläimet)
      |  |?- Cycliophora Funch & Kristensen, 1995 (“Wheel Wearers”)
      |  `?- Mesozoa [see Basal Metazoa]
      `--+?-o †Cribricyathea
         |  |-- †Conoidocyathida
         |  |-- †Cribricyathida
         |  `-- †Vologdinophyllida
         |?-o †Machaeridia
         |  |-- †Hercollepadida
         |  `-- †Turrilepadida
         |--o Ecdysozoa Aguinaldo, Turbeville, Linford, Rivera, Garey, Raff & Lake, 1997 (skinshedders; nahanluojat)
         |  |-- †Facivermis yunnanicus (transional for between pripulid/palaeoscolecid worms and lobopods??)
         |  |--o Arthropodomorpha (arthropods and related animals; niveljalkaismaiset eläimet)
         |  |  |?-o †Sprigginida
         |  |  |  |?- †Bomakellia kelleri Fedonkin 1985 [a rangeomorph frond]
         |  |  |  `-- †Spriggina floundersi Glaessner, 1958 [also a rangeomorph frond?]
         |  |  `--o Aiolopoda Hou & Bergström, 1996 [incl. Arthropoda von Siebold, 1848][Panarthropoda] (water bears, velvet worms, anomalocarids and arthropods; karhukaiset, käsnäjalkaiset, anomalocaridit ja niveljalkaiset)
         |  |     |-- Tardigrada Spallanzani, 1777 (water bears; karhukaiset) [considered as Aschelmithes by Hou & Bergström (1997)]
         |  |     `==o Loboboda Snodgrass, 1938 [Onychophora sensu lato] [Microdictyoniida; Protonychophora Hutchinson, 1930] (lobopods, velvet worms; käsnäjalkaiset) [paraphyletic grade of earliest arthropods]
         |  |        |-- Onychophora Grube, 1853 [Euonychophora Hutchinson, 1930] (modern lobopods, velvet worms; nykykäsnäjalkaiset)
         |  |        `--+-- †Radiodonta Collins, 1996 (anomalocaritids and relatives)
         |  |           `-- Euarthropoda [Arthropoda sensu stricto] [Schizoramia sensu Hou & Bergström, 1997]
         |  `--o Scalidophora sensu Dong, Donoghue, Cheng & Liu, 2004
         |     |--+-- †Ancalagon minor (Walcott, 1911) Conway Morris, 1977 [Ottoia minor Walcott, 1911] [Ancalagonidae Conway Morris, 1977]
         |     |  `-- †Fieldia lanceolata Walcott, 1912 [Fieldiidae]
         |     `--+--o †Markuelia Val´kov, 1983 [Middle and Late Cambrian protostome embryos]
         |        |  |-- †M. hunanensis Dong & Donoghue, in Dong, Donoghue, Cheng & Liu, 2004
         |        |  `-- †M. secunda Val´kov, 1983
         |        `--o Priapulida Delage & Hérouard, 1897 (non Théel, 1906?) sensu lato [Priapuloidea] [incl. Palaeoscolecida Conway Morris & Robison, 1986 sensu Dong, Donoghue, Cheng & Liu, 2004] (priapulid worms; makkaramadot)
         |           |==Palaeoscolecida Conway Morris & Robison, 1986 [paraphyletic basal Scalidophorans?]
         |           `--+-- Kinorhyncha Reinhard, 1881 (mud dragons; okakaulamadot)
         |              `--+-- Priapulida Delage & Hérouard, 1897 (non Théel, 1906?) (s.s.) [Priapuloidea] (priapulid worms, penis worms; makkaramadot)
         |                 `-- Loricifera Kristensen, 1983 (Brush heads)
         `--o Coeloscleritophora Bengtson & Missarzhevsky, 1981 [paraphyletic]
            `--o Lophotrochozoa Halanych, Bacheller, Liva, Aguinaldo,, Hillis & Lake, 1995
               |--o “Molluscamorpha” [Mollusca sensu lato]
               |  |-- †Kimberella quadrata (Glaessner & Wade, 1966) Wade, 1972 [Kimberia quadrata Glaessner & Wade, 1966]
               |  `--+-- †Odontogriphus omalus Conway Morris, 1976 [Odontogriphidae Conway Morris, 1976]
               |     |?=Halwaxida Conway Morris & Caron, 2007 [incl. Siphogonuchitida; Halkieriida; Wiwaxiida; ?Sachitida; ?Chancelloriida] [paraphyletic basal Aculiferas sensu Vinther, 2015]
               |     `--+?- †Tullimonstrum gregarium Richardson, Jr., 1966  [Tullimonsterida] [has been reidentified as hyperoartian vertebrate by McCoy et al. (2016)]
               |        `--+-- Mollusca Cuvier 1795 (molluscs; nilviäiset)
               |           `--+-- †Hyolitha Marek, 1963 (hyoliths, hyoliitit)
               |              `-- Sipuncula Rafinesque, 1814 [Sipunculoidea] (sipunculid worms, peanut worms; ruiskumadot)
               `--+--o “Annelidamorpha”
                  |  |-- †Canadia spinosa Walcott, 1911
                  |  |--+-- †Amiskwia sagittiformis Walcott,1911 [Amiskwiida]
                  |  |  `-- Nemertea Schultze, 1851 [Nemertinea] (ribbon worms, proboscis worms; limamadot)
                  |  `--o Annelida sensu lato
                  |     |-- Lobatocerebrum psammicola Rieger, 1980 [Lobatocerebrida; Lobatocerebridae Rieger, 1980]
                  |     `--+--+?- †Banffia constricta [Banffozoa: Banffida: Banffidae]
                  |        |  `-- Echiura Newby, 1940 [Echiuroidea] (spoon worms; tähtimadot)
                  |        `-- Annelida Lamarck, 1809 [incl. Vestimentifera & Pogonophora sensu Rouse, 2001] (annelids, segmented worms; nivelmadot)
                  `--o Lophophorata
                     |-- Phoronida Hatschek, 1888 (phoronids, horseshoe worms; tupsumadot)
                     `--+==Tommotiida Missarzhevsky, 1970 (tommotians)
                        `-- Brachiopoda Duméril, 1806 (brachiopods, lampshells; lonkerojalkaiset)

After Aguinaldo, Turbeville, Linford, Rivera, Garey, Raff & Lake, 1997 and Dong, Donoghue, Cheng & Liu, 2004

<==o Protostoma
   |?- Myzostomida
   |?- Annelida
   |?- Mollusca
   |?- Sipuncula
   |--o Lophotrochozoa
   |  |?- Mesozoa
   |  |?- Entoprocta
   |  |?- Bryozoa [Ectoprocta]
   |  |?- Cycliophora
   |  |?- Gastrotricha
   |  |?- Rotifera
   |  `?- Acanthocephala
   `--o Ecdysozoa
      |?- Chaetognatha
      |--o Introverta
      |  |--+-- Nemata
      |  |  `-- Nematomorpha
      |  `--o Scelidophora
      |     |--+-- †Ancalagon
      |     |  `-- †Fieldia
      |     `--+--o †Markuelia
      |        |  |-- †M. hunanensis
      |        |  `-- †M. secunda
      |        `--+-- Kinorhyncha
      |           `--+-- Loricifera
      |              `-- Priapulida
      `--o Euarthropoda
         |-- Loboboda
         |-- Tardigrada
         |-- Crustaceomorpha
         |-- Insecta
         `--+-- Myriapoda
            `-- Chelicerata

Reference(s):

  • Aguinaldo, A. M. A., Turbeville, J. M., Linford, L. S., Rivera, M. C., Garey, J. R., Raff, R. A. & Lake, J. A., 1997: Evidence for a clade of nematodes, arthropods and other moulding animals.
    –Nature: Vol. 387, 29 May, pp. 489 - 493
  • Dong, X.-p., Donoghue, P. C. J., Cheng, H. & Liu, J.-b., 2004: Fossil embryos from the Middle and Late Cambrian period of Hunan, south China.
    –Nature: Vol. 427, pp. 237-240

After Winnepennickyx et al., 1998

<==o Protostoma
   |?- Mesozoa
   |?- Sipuncula
   |?- Myzostomida
   |?- Loricifera
   |?- Chaetognatha
   |?- Kinorhyncha
   |?- Nematomorpha
   |-- Mollusca
   |-- Annelida
   |--o Lophophorata
   |  |-- Phoronida
   |  `-- Brachiopoda
   |-- Entoprocta
   |-- Bryozoa [[Ectoprocta]
   |-- Gastrotricha
   |--+?- Cycliophora
   |  `--+?- Rotifera
   |     `?- Acanthocephala
   |-- Lobopoda
   |-- Nemata
   `--+-- Priapulida
      `--+?- Tardigrada
         |?- Myriapoda
         |?- Chelicerata
         `--+-- Crustaceomorpha
             `-- Insecta

Note:

This tree is from molecular phylogenetic study made by Winnepennickyx, Backeljau & Kristensen (1998). This doesn't agree with a study by Aguinaldo et al. (1997), although it too contains a clade made by priapulids and arthropods. Aguinaldo et al. also contains clade made by phoronids and brachiopods.

Several Protostome-phylums were apparently untreated, I have placed them here too, marked as unsure relationship. [ |?- ]


Reference(s):

  • Winnepennickyx, B. M. H., Backeljau, T. & Kristensen, R. M., 1998: Relationships of a new phylum Cycliophora.
    –Nature: Vol. 393, 18 June, pp. 636-638

After Zrzavý et al., 2001, with fossil phyla from Conway Morris, 1993

<==o PROTOSTOMA
   |--o Aschelmithes
   |  |-- Xenoturbellida
   |  |-- Platyhelmithes
   |  `--o Acanthognatha
   |     |--o Neotrichozoa
   |     |  |-- Gastrotricha
   |     |  `-- Gnathostomulida
   |     `--o Prosomastigozoa
   |        |-- Myzostomida
   |        `--+-- Cycliophora
   |           `--o Syndermata
   |              `==o Rotifera
   |                 |-- Monogononta
   |                 `--o Lemniscea
   |                    |-- Bdelloida
   |                    `--+-- Seisonida
   |                       `-- Acanthocephala
   `--+--o Arthropodomorpha
      |  |-- Cephalorhyncha
      |  |--+-- Chaetognatha
      |  |  `--+-- Nematomorpha
      |  |     `-- Nemata
      |  `--+--o †Sprigginida
      |     |  |?- †Bomakellia
      |     |  `-- †Spriggina floundersi
      |     |?- Tardigrada
      |     `-- Arthropoda
      `--+-- Entoprocta
         |-- Bryozoa [Ectoprocta]
         |--+-- †Chancelloriida
         |  `-- †Sachitida
         `--+-- †Siphogunuchitida
            `--+--o †Halkieriida
               |  `-- †Halkeria evangelista
               `--+--o “Molluscamorpha”
                  |  `--+?- †“Tullimonsterida”
                  |     `--+-- Mollusca
                  |        `--+-- †Hyolitha
                  |           `-- Sipuncula
                  `--+--o “Annelidamorpha”
                     |  |?- †Wiwaxia corrugata
                     |  `--+?- †Canadia spinosa
                     |     |--+-- †Amiskwia sagittiformis
                     |     |  `-- Nemertea
                     |     `--+-- Lobatocerebrida
                     |        `--+--+-- †Banffida
                     |           |  `-- Echiura
                     |           `-- Annelida
                     `--+--o Lophophorata
                        |  |-- Phoronida
                        |  `--+-- †Tommotiida
                        |     `-- Brachiopoda
                        `--+-- †Palaeoscolecida
                           `--+-- Kinorhyncha
                              `--+-- Loricifera
                                 `-- Priapulida

Reference(s):

  • Zrzavý, J., Hypsa, V. & Tietz, D. F., 2001: Myzostomida are not annelids: Molecular and morphological support for a clade of animals with anterior sperm flagella.
    –Gladistics: Vol. 17, #2, pp. 170-198

After Brusca & Brusca, 2002

<==o Bilateralia
   |--o Deuterostomia
   |  |-- Chaetognatha
   |  |-- Ectoprocta
   |  |-- Phoronida
   |  |-- Brachiopoda
   |  |-- Echinodermata
   |  `--+--o Hemichordata
   |     |  |-- Pterobranchia
   |     |  `-- Enteropneusta
   |     `--o Chordata
   |        |-- Urochordata
   |        `--+-- Cephalochordata
   |           `-- Vertebrata
   `--o Protostomia
      |-- Platyhelmithes
      `--+-- Nemertea
         |--+-- Sipuncula
         |  `-- Mollusca
         |-- Echiura
         |--o Articulata
         |  |-- Annelida
         |  `--+-- Onychophora
         |     `--+-- Tardigrada
         |        `-- Arthropoda
         |-- Gnathostomulida
         |--+-- Entoprocta
         |  `-- Cycliophora
         |--+-- Rotifera
         |  `-- Acanthocephala
         `--o Cycloneuralia
            |-- Gastrotricha
            |-- Nemata
            |-- Nematomorpha
            `--+-- Priapulida
               `--+-- Kinorhyncha
                  `-- Loricifera

Reference(s):

  • Brusca, R., C. & Brusca, G., J., 2002: Invertebrates.
    –Sinauer Associates Inc., Sunderland, Massachusetts, 2002, xix-936

After Paps, Bagañà & Riutort, 2009

<==o Protostoma
   |--o Ecdysozoa (nahanluojat)
   |  |?- Chaetognatha
   |  |--+-- Nematomorpha
   |  |  `-- Nemata
   |  `?-+--+-- Kinorhyncha
   |     |  `-- Priapulida
   |     `--+-- Tardigrada
   |        `-- Arthropoda
   `--o Lophotrochozoa sensu lato
      |-- Gnathostomulida
      `--+-- Gastrotricha
         `--+--o Gnathifera
            |  |-- Limnognathia maerski [Micrognathozoa]
            |  `--+== Rotifera
            |     `-- Acanthocephala
            `--o Lophotrochozoa sensu stricto
               |--+-- Platyhelmithes
               |  `--+-- Bryozoa [Ectoprocta]
               |     `--+-- Cycliophora
               |        `-- Entoprocta
               `--+-- Nemertea
                  `--+--+-- Echiura
                     |  `--+-- Sipuncula
                     |     `-- Annelida
                     `--+-- Mollusca
                        `--o Brachiozoa
                           |-- Phoronida
                           `-- Brachiopoda

Reference(s):

  • Paps, J., Bagañà, J. & Riutort, M., 2009: Lophotrochozoa internal phylogeny: new insights from an up-to-date analysis of nuclear ribosomal genes.
    –Proceedings of the Royal Society of London: Biological series: Vol. 276, pp. 1245-1254 [doi: 10.1098/rspb.2008.1574]

Reference(s):